H. Mai et al.: Paleocene coccospheres from DSDP Leg 39 .... ... , p. 21 - 29.

Journal of Nannoplankton Research, 20, I , 1998.

PALEOCENE COCCOSPHERES FROM DSDP LEG 39, SITE 356,
SAO PAULO PLATEAU, S ATLANTIC OCEAN
Hartmut Mai*, Tania Hildebrand-Habel*, Katharina von Sa/is Perch-Nielsen** & Helmut Willems**Universittit Bremen, Fachbereich 5- Geowissenschaften, Klagenfurter Strasse, D-28359 Bremen, Germany;**Geologisches Jnstitut, ETH-Zentrum, CH-8092, Zurich. Switzerland
jf,

l "!

•

Abstract: Numerous coccospheres representing the two extant families , Coccolithaceae and Prinsiaceae,
were observed in 12 Paleocene (NP2 through NP6) samples from DSDP Site 356, and are briefly described(Calcidiscus? sp., Chiasmolithus edentulus, Coccolithus pelagicus, Cruciplaco/ithus primus ,Crucip/aco lithu s tenuis, Cmciplacolithus sp., Cyclagelosphaera alta, Ericsonia subpertusa, Ericsonia?
sp., Futyania petalosa. Markalius c( Markalius inversus, Prinsius dimorphosus, S ul/ivania consueta,
Sullivania sp., Toweius eminens, Toweius pertusus). Upper Maastrichtian (CC26) and lowermost Danian(NPI) sediments furnished only coccospheres of Watznaueria barnesae, whereas samples covering thezones from NP2 to NP6 e xhibit ed an increased numb er of different species. The exce llent state ofpreservation of coccoliths from samples 39-356-29-1. 83-85cm (NP2) and 39-356-21-2, 122-123 cm(NP5) may be due to rapid deposition, possi bly within fecal pellets, and minor diageneti c effects. We
conclude that the presence of intact fossil coccospheres occurs if a sediment is rich in well-preserved
coccoliths; their observation then depends on an adequ ate preparation met hod and the tim e spent withthe scanning electron microscope.

Introduction
Fossil coccoliths are not usually found forming
coccospheres but are distributed as single platelets on theslide prepared for investigation by light rAicroscope (LM)or scanning electron microscope (SEM) . Some fossil
coccospheres are, however, occasionally found andillustrated. Watznaueria barnesae, numerous species ofBiscutum , Coccolithus pe/agicus , and various
representatives of small- and medium-sized Prinsiaceae are
the most common fonns reported as coccospheres.
The intensive SEM study of the Kif boundary claysof Geulhemmerberg in The Netherlands (Mai et al., 1997)
resulted in the illustration and description of more than 30
Maastrichtian and lowennost Danian coccosphere species,
many ofthem well preserved.
While investigating the Kif boundary interval of
DSDP Leg 39, Site 356 for calcareous dinoflagellate cysts, awell-preserved coccosphere of Markalius cf. Af. inversus
was found in sample 39-356-21-2, 122-l23cm (NP5 ofMartini, 1971 ). The subsequent discoveries of coccospheresof Futyania peta/osa in sample 39-356-29-l, 83-85cm (NP2of Martini, 1971) triggered a major effort to find additional
coccospheres. Perch-Nielsen (1977) had only illustrated
coccospheres of Prinsius dimorphosus from sample 39356-28-3, 70(NP3ofMartini, 1971).

Systematic palaeontology
The genera are arranged in alphabetical order.
Genus Calcidiscus Kamptner, 1950

Calcidiscus? Sll.
Plate 1, Figure 1
The spherical coccosphere measures 21 J.Ull in diameter andconsists of approximately 20 circular to elliptical coccolithsof equal size resembling those of the Neogene and Recent
genus Calcidiscus. The coccoliths overlap considerably.
The distal shields consist of approximately 38 elements andare larger than the proximal shields. The curved elements
imbricate dextrally and leave only a small, conical central
depression.
Genus Chiasmolithus Hay et al, 1966 emend. Varol ,1992

Chiasmolithus edentulus van Heck & Prius, 1987
Plate I, Figures 2, 3, 6
The coccospheres are 12J.UU to l8J.Un in diameter, sphericalto slightly ovoid and consist of ten to 16 elliptical,
overlapping coccoliths of about equal size. Each coccolithis surrounded by five or six other coccoliths. The central
cross is similar in all visible coccoliths ofthe coccospheres.

Material and methods
The samples studied were collected from DSDP Leg 39 at
Site 356 on the Sao Paulo Plateau in the South Atlantic at
28°17.22 ' S, 41005 .28'W, 950km offshore from Brazil at awater-depth of3175m. The sediments mainly consist of purenannofossil and foraminiferal chalks with varying amountsof clay. The ages of the 12 samples studied range fromlatest Maastrichtian CC25 to Late Paleocene NP6 (PerchNielsen, 1977). The samples were prepared following themethod described by Mai et al. ( 1997), which is lessdestructive for coccospheres than the methods usually used.
The total time for SEM analysis in search of coccospheres
amounted to more than 500 hours.
The negatives referred to in the plate descriptions
are stored in the Fachgebiet Historische Geologie/
Palaontologie, Fachbereich 5, Bremen University.

Genus Coccolithus Schwartz, 1894

Coccolithus pelagicus (Wallich, 1877) Schiller, 1930
Plate l , Figures 4, 5
The coccospheres are lOJ.UU to 18J.UU in diameter, sphericaland resemble the modem coccospheres ofthe species. The
number of coccoliths varies from nine to 14 in the spheres
illustrated here, a number comparable to that of Recent
coccospheres. The equally-sized coccoliths overlap
considerably and each coccolith is surrounded by five othercoccoliths.

21

Journal of Nannopl ankt on Research, 20. I , 1998.

H . M01 e 1 a l.: Paleocene coccospheres from DSDP Leg 39 ...... , p. 2 1 - 29 .

Genus Cruciplacolithus Hay & Mohler in Hay et aL
1967

coccoliths of equal size. As each coccolith is surroundedby four coccoliths. the coccospherc is cube-shaped. It
measures approximately IOJ.!m . The illustrations of E.
subpertusa in Pcrch-Nielsen ( 1977) and Mai et al. ( 1997)
also show coccosphcrcs with only six coccoliths. The
coccosphcre illustrated as Ericsonia cf. F. subpertusa in
Perch-Nielsen ( 1971) from the Lower Eocene of Denmark.
however. has approximately 12 coccoliths, each surroundedby five others.

Cruciplacolithusprimus Perch-Niclsen 1977
Plate 2, Figures 1-3
The coccosphcrcs are spherical to slightly ovoid withdiameters of between 6J.iln and 16~un , and consist of ca .1218 coccoliths which show variation in overall size. Eachcoccolith is surrounded by five or six other coccoliths. The
number of elements fonning the rims of the coccoliths seems
to be similar within coccoliths of the same coccosphcrc.
and increases with the size oft he coccolith.

Ericsonia? sp.
Plate 3, Figure 5
The subspherical coccospherc consists of approximately
14 elliptical coccoliths, each of them surrounded by fiveothers. The sphere measures I5J.!m in diameter. and thecoccoliths are of equal size. The distal shields arc relatively
narrow and steep and the crown is formed by elements ofthe tube-cycle. The central opening is elliptical. The formshows features typical of Ericsonia, but the central-area
elements seem to be arranged radially, as in Coccolithus,
rather than imbricating, as in E. subpertusa.

Cruciplacolithus tenuis (Stradner, 1961) Hay &Mohler in Hay et al., 1967
Plate 2. Figure 6
The coccosphere measures 19~un . it is spherical and consistsof approximately 13 interlocking coccoliths of similar size.
Each coccolith is surrounded by five other coccoliths. The
centrcll cross is clearly a-dally-aligned and all four anns bc-1I
tenninal feet.

Cruciplacolithus sp.
Plate 4, Figure 6

Genus Futyania Varol, 1989

Futyania petalosa (EIIis & Lohman, 1973) Varol, 1989
Plate 3, Figures 1-3

The subcircular coccoliths of this subsphcrical coccosphercfeature a central structure consisting of a cross with aslightly rotated long axis . The coccospherc is constructedof approximately 14 coccoliths . Single coccoliths arc
surrounded by five or six others. It measures 11 J.ill1 .

The coccosphcres of F petalosa are spherical. IOJ.!m to
I2J.!m in diameter. and consist of approximately 80 small ,elliptical coccoliths. Most of them are surrounded by six
others. sometimes by seven. The presence of the petaloid
distal elements seems to depend on the state of preservation.

Genus Cyclagelo.\ phaera Noel, 1965

Cyclagelosphaera alta Perch-Niclscn, 1979
Plate 2, Figures 4, 5Genus Markalius Bramlctte & Martini, 1964

Markalius cf. M. inversus (Deflandre in Dcflandre &Fcrt, 19S4) Bramlette & Martini, 1964
Plate 3, Figures 6-9

The coccospheres are spherical and 11 J.Un to I5J.!m indiameter. They consist of 12 coccoliths of equal size. Eachcoccolith is surrounded by five other coccoliths. The
number of radial to tangential ridges forming the central
cone varies on the coccosphcre from a minimum of five toa maximum of seven. In the two coccosphercs illustrated
here. the central cone has a pentagonal to polygonal central
depression .

The spherical coccosphcrcs are constructed ofapproximately 12 more-or-less circular coccoliths of equalsize. The coccospheres are 17 J.ill1 to 19 J.ilTI in diameter. Eachcoccolith is surrounded by five other coccoliths. The
central areas of the coccoliths are fonncd by several layersof approximately radially-arranged. polygonal elements
which do not reach the crown of the distal shield.

Genus Ericsonia Black, 1964
The genus Ericsonia is here retained for the forms whichfeature a relatively narrow distal shield, although many
authors include these within Coccolithus. The elements ofthe tube form a crown over the distal shield and areoverlapping in a clockwise direction around a narrow
central opening. In Coccolithus, the distal shield is wider
and the elements of the tube-cycle are arranged more-orless radiaJiy.

Genus Prinsius Hay & Mohler, 1967

Prinsius tfimorphosus (Perch-Nielsen, 1969) PerchNielsen, 1977
Plate 4, Figure I
The coccosphere measures lOJ.!m ;~ diameter and iscomparable with those illustrated in the existing literature.
Each coccolith is surrounded by six other coccoliths, which
are equal in size. The total number of coccoliths on the
coccosphere is approximately 20.

Ericsonia subpertusa Hay & Mohler, 1967
Plate 3, Figure 4
The coccosphere is formed from only six subcircular22

= - - - - - - - -- -- - - - -H. M01 er al.: Paleocene coccosphercs from DSDP Leg 39 ... .. ... p. 21 • 29.
' V

Genus Su/livania Varol, 1992

Sullivania consueta (Bramlette & Sullivan, 1961)
Varol, 1992
Plate 4. Figure 2

I
This large coccosphcrc (2l!Jm) is subsphcrical and consistsof 16 coccoliths of equal size. Each coccolitJt is surroundedby either five or six other coccoliths. The fonn illustrated ischaracterised by a central diagonal cross with arms ofapproximately equal length . The crown of the coccolith isformed by the distal shield. a characteristic feature of thegenus Sullivania . The structure of the arms is not clearlyvisible, but there seems to be no indication of parallel.
longitudinal laths, as is typical of forms belonging to thegenus Chiasmolithus.

Sullivania SJ).
Plate 4, Figures 3-5
The coccospheres measure II!Jm to I51Jm . They arespherical and contain seven to 12 overlapping coccolithsof equal size. The central areas of the coccoliths are relatively
small and spanned by a central diagonal cross. Each arm of
the central structure connects to a foot tJtat lines the central
opening. Each coccolith is surrounded by five othercoccoliths, both on coccospheres with only sevencoccoliths and those with more than ten coccoliths.

<knus Toweius Hay & Mohler, 1967

Toweius eminens (Bramlette & Sullivan, 1961) PerchNielsen, 1971
Plate 5, Figure I
As the illustrated coccosphere of l31Jm in diameter isprobably not completely preserved (some coccoliths
missing?) the shape cannot be exactly determined. Eachcoccolith exhibits four central openings, characteristic of
this species.

Journal ofannoplankton Research, 20, I, 1998.

treatment (Mai et al .. 1997). Coccosphercs can also be
studied by direct observation of rock surfaces (e.g. Bown.
1993, and others).
-The time for SEM analysis must be greatly increased
compared to the normal time spent looking for dispersed
coccoliths.
In the lower part of the section studied , thecoccospherc assemblages are dominated by relatively
poorly preserved Watznaueria barnesae, not illustrated
here. both in the uppermost Cretaceous (CC26) and in thelowem1ost Tertiary (NP I) samples (sec Appendix).
The coccosphercs from sample 39-356-29-1. 83-85cm(NP 2) are. like the dispersed coccolitJts in this sample, very
well-preserved and include Coccolithus pelagicus,
Futyania petalosa and Prinsius dimorphosus. In theinterval representing NP3 and NP4, only relatively poorlypreserved coccosphcres of C. pelagicus and ruciplacolithus prim us were encountered. The sample richestin coccospheres (39-356-21-2, 122-123cm) belongs to theupper part of NP5 and yielded 13 species represented bywell-preserved coccospheres. In the samples above this,
only coccosphercs of . pelagicus were found .

Coccospherc morphology:
An attempt was made to determine the number of coccolitlts surrounding each coccolith in the various species.
The preliminary results indicate that many species tend tohave five surrounding coccoliths, so that the resulting
shapes are pentagonal dodecahedra . Some others usually
have six in some the number varies, and some fonns seemto have only four coccoliths surrounding each coccolith.
The latter leads to a more-or-less cubic coccosphcre, as
observed in Ericsonia subpertusa .
It may or may not be a coincidence thatMarkalius
cf. M. inversus and Cyclagelosphaera alto, both forminga pentagonal dodecahedron-like structure. belong to genera
that survived the KIT boundary cvcnt(s) . Also ,enToweius pertusus (Sullivan, 1965) Romein, 1979
Plate 5 Figures 2-5
The coccospheres here assigned to T pertusus are from
6~tm to 9~tm in diameter and arc subsphericaJ. They consist
often to 12 elliptical. interlocking coccoliths. Each coccolitlt
is surrounded by four to five other coccoliths of about
equal size.
Results
Presence of coccOSJ>hcrcs:
We suggest that the finding of intact coccospheres (otherthan Hatznaueria barnesae) depends mainly on thefollo\\i ng condi lions:-The sediment inYestigated must be rich in coccoliths.
lithsm t be'' 11-preserYed.
lr::t!Jnn
cti\ forbe n-d....... ·'---· uf-~:-h . ~ -~ :--..:=::~....;.ci:al _•....:iiu:- of-......

~==(C) Coccolithaceae(P) Prinsiaceae(,!)~=~ ~~~
0!-<CII:: 0~-

cc:S~ CCj:)..J~oo

18~
Uc.: ~ccuc.:U(FAMILY)
SHAPE OF
0"'1
SPECIESCOCCOSPHERE
u~ z~8spherical(C) Calcidiscus ? sp.
20
5-6
spherical
10-16 5-6(C) Ch. edentulus
spherical
9-14(C) C. pelagicus
5(C) Cr. primus[Spherical to ovoid
12-18 5-6(C) Cr. tenuis
spherical
135(C) Cruciplacolithus sp. :spherical
14
5-6
pentagonal dodecahedron 12(C?) Cy. alta
5(C) E. subpertusa
cubic
4
6
ovoid
(C) Ericsonia sp.
145(P) F. petalosaspherical
6-780(C?) M. cf. M. inversus pentagonal dodecahedron 12
5(P) P. dimorphosus
spherical
205(C) S. consueta
spherical
16
5-6
spherical(C) Sullivania sp.
7- 12
5(P) T. eminens
spherical to cubic?
>6>3(Pl T. pertusus
spherical to cubic?
10-12 4-5

Ta* I : ~ o( ;:-;"'--~ ~...,._._ '~""T o1 - .,_..........,.. aad m:~'""'crIJournal of Nannoplankton Research, 20, I , 1998.

H . Mai et al.: Paleocene coccospheres from DSDP Leg 39 ....... , p. 21 - 29.

Braarudosphaera and Goniolithus, two other survivors

Mai, H ., von' Sa lis Perch-Nielsen, K .. Willems, H. & Romein , T.

of the K/T boundary event(s) , form pentagonal
dodecahedra, but this is due to the pentagonal outlines ofsingle coccoliths.
As in Recent coccolithophorid species. the numberof coccoliths varies on fossil coccospheres. While somespecies, like Markalius cf. M. inversus and Cyc!agelosphaera a/ta, show a constant number of coccoliths on
the sphere, others include a certain variation of numbers,
like Futyania petalosa (Table l ).

1997. Fossi l coccospheres from the K IT boundary section
from Geulhemmerberg, The Netherlands. Micropale011tology.

Conclusions
We conclude that more coccospheres might be found thanare commonly reported if the suggestions set out above
are followed. The fact that even the 500 hours spent on the
SEM did not result in the discovery of somet11ing similar to
coccospheres oft11e systematically ambiguous fasciculiths,
sphenoliths or heliolit11s does not necessarily indicate t11atthere are no coccospheres of these taxa, but maybe due totheir shape, they do not build preservable coccospheres(not interlocking).
Species which occur as coccospheres belong to only
two families, the Coccoli thaceae and the Prinsiaceae. They
have living representatives and dominate calcareous
nannoplankton assemblages of the upper photic zone. In
the present day the Prinsiaceae are represented byEmiliania hux/eyi and species of Gephyrocapsa.

43(3): 281-302.
Martini, E. 1971. Standard Tertiary and Quaternary calcar~ous
nannoplankton zonation . /n: A. Farinacci (Ed .). Proc. If lnt.
Plankt. Conf. , Roma. 1970. Tecnoscienza, 2: 739-785 .
Perch-Nielsen , K . 1971 . Elektronenmikroskopische
Untersuchungen an Coccolithen und verwandten Formen a us
dem Eozan von Danemark. Bioi. Skr. Dan. Vid. Selsk .• 18(3):1-76.
Perch-Nielsen , K. 1977 . Albian to Pleistocene ca lca reou s
nannofossils from the western South Atlantic. /JWSDP, 39:
699-823 .
Perch-Nielsen, K . 1985. Cenozoic calcareous nannofossils . In :II.M. Bolli. J.B. Saunders & K. Pcrch-Nielsen (Eds). Plankton
Stratigraphy. Cambridge University Press: 427-554.
Varol, 0 . 1989. Paleocene calca reous nannofossil biostratigraphy.
In : J.A. Crux & S.E. van Heck (Eds). Nannofossils and theirapplications. Ellis Horwood: 267-310.
Varol, 0 . 1992. Sullivania : a new genus ofPalcogene coccoliths.

J.Micropalaeont., 11 : 141-150.

AJ)J)Cndix: Investigated samJ>Ics

Acknowledgements
The DSDP/ODP is thanked for providing the authors withmaterial. Financial support was given by Project Wi 725/ I 0
oft11e German Science Foundation (Deutsche Forschungsgemeinschaft). Many thanks to Jackie Burnett and Paul
Bown for their constructive and critical review of themanuscript.

~
>
~

"'

rzl

rzl

~

....rzl
~
s
u

~

1:1.

<
~

"'<t=

0
rzl 5:::~
COCCOSPHERES
..J m
u
m
NP6 C. pelagicus
39 356 17-4 75-77
39 356 18-3 130-133 NP5 C. pe/agicus
NP5 C. pelagicus
39 356 19-2 74-75
39 356 21 -2 122- 123 NP5 Most of taxa described
39 356 22-4 77-78 . NP5 C. pelagicus, C. primus
39 356 24-2 77-78
NP5 C. pelagicus, C. primus
NP4 C. pelagicus, C. primus
39 356 25-4 60-62
NP4 C. pelagicus, C. primus
39 356 26-5 77-78
NP3 C. pe/agicus, C. primus
39 356 27-6 75-76
NP2 F. petalosa, C. pelagicus, P. dimorphosus
39 356 29- 1 83-85
39 356 29-2 131 - 133 NP1 W. barnesae
39 356 29-3 106-108 CC26 W. barnesae

Refer·ences
Further taxonomic references can be found in Perch-Nielsen(1985).
Bown, P.R . 1993. New holococcoliths from the Toarcian-Aalenian(Jurassic) of northern Germany. Senckenbergiana lethaea.
73: 407-419.
van Heck, S .E . & Prins, B . 1987.Arefined nannoplanJ...1on zonation
for the Danian of the central North Sea . A bh. Geol. B.-A ,Wien , 39: 285-303.

'

24

H. Mai et al.: Paleocene coccospheres from DSDP Leg 39 ... .... , p. 21 - 29.

Journal of Nannoplankton Research, 20, 1, 1998.

PLATE 1

Chiasmolithus edentulus

Chiasmolithus edentulus

Calcidiscus? sp.

Coccolithus pelagicus

Coccolithus pelagicus

Chiasmolithus edentulus

PLATE 1: Fig. 1: Calcidiscus? sp .. Sample 39-356-21-2, l22-123cm (NP5); Neg.# 867/32. Figs 2,3, 6: Chiasnwlithus
edentulus. Sample 39-356-21-2, 122-123cm (NP5); 2: Neg.# 867/32; 3: Neg.# 867/4; 6: Neg.# 867/17. Figs 4, 5:
Coccolithus pelagicus. Sample 39-3 56-21 -2, 122-123cm (NP5); 4: Neg.# 854/12; 5: Neg.# 854/24.

25

Journal of Nannopl ankton Research, 20, I, 199&.

H. Mai et al.: Paleocene coccospheres from DSDP Leg 39 ....... , p. 2 1 - 29.

PLATE 2

Cruciplacolithus primus

Cruciplacolithus primus

Cruciplacolithus primus

Cyc/agelosphaera alta

Cyclage/osphaera a/ta

Cruciplacolithus tenuis

PLATE 2: Figs 1-3: Cruciplacolithus primus. Sample 39-356-21-2, 122-123cm (NP5); 1: Neg.# 85417; 2: Neg.#
854/32; 3: Neg.# 854/14. Figs 4, 5: Cyclagelosphaera alta. Sample 39-356-21-2, 122-123cm (NP5); 4: Neg.# 853/
27; 5: Neg.# 853/8. Fig. 6: Cruciplacolithus tenuis. Sample 39-356-21-2, 122-123cm (NP5); Neg.# 854/28

26

H. Mai et al. : Paleocene coccospheres from DSDP Leg 39 ....... , p. 21 - 29.

Journal ofannoplankton Research, 20, I , 1998.

PLATE 3

Futyania petalosa

Futyania petalosa

Futyania petalosa

Ericsonia subpertusa

Markalius cf. M. inversus

Markalius cf. M. inversus

Ericsonia? sp.

Markalius cf. M. inversus

Markalius inversus

PLATE 3: Figs 1-3: Futyania petalosa. Sample 39-356-29-1 , 83-85cm (NP2); 1: Neg.# 854/16; 2: Neg.# 854/9;
3: Neg.# 854/24. Fig. 4: Ericsonia subpertusa. Sample 39-356-21-2, 122-123cm (NP5); Neg.# 853/2. Fig. 5:
Ericsonia? sp .. Sample 39-356-21-2, 122-l23cm (NP5); Neg.# 853/3. Figs 6-9: Markalius cf. M. inversus. Sample
39-356-21-2, l22-123cm (NP5); 6: Neg.# 853/5 ; 7: Neg.# 853/6; 8: Neg.# 853/12; 9: Neg.# 853/13 .

27

Journal of Nannoplankton ~esearch, 20, I , 1998.

H. Mai et al.: Paleocene coccospheres from DSDP Leg 39 ..... .., p. 21 - 29.

PLATE 4

Prinsius dimorphosus

Sul/ivania consueta

Sul/ivania sp.

Sul/ivania sp.

Sullivania sp.

Cruciplacolithus sp.

PLATE 4: Fig. 1: Prinsius dimorphosus. Sample 39-356-29-1 , 83-85cm (NP2); Neg.# 854/3 . Fig. 2: Sullivania
consueta. Sample 39-356-21-2, 122-123cm (NP5); Neg.# 854/5. Figs 3-5: Sullivania sp .. Sample 39-356-21-2,
122-123cm (NP5); 3: Neg.# 854/28; 4: Neg.# 854/24; 5: Neg.# 854/l9. Fig. 6: Cruciplacolithus sp.. Sample 39356-21-2, 122-123cm (NP5); Neg.# 854/4.

28

H. Mai et al.: Paleocene coccospheres from DSDP Leg 39 ....... , p. 21 - 29.

Joumal of Nannoplankton Research, 20, I , 1998.

PLATE 5Toweius pertusus

Toweius pertusus

Toweius pertusus

Sediment {NPS)

Toweius pe_rtusus

PLATE 5: Fig. 1: Toweius eminens. Sample 39-356-29-1 , 83-85cm {NP2); Neg.# 853/19 . Figs 2-5: Toweius
pertusus. Sample 39-356-21-2, 122-123cm (NP5); 2: Neg.# 853/22; 3: Neg.# 853/20; 4: Neg.# 853/9; 5: Neg.#
853/16. Unnumbered: Sediment of NP5 at low magnification.

29